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Trade-Offs Between Burrowing and Biting Force in Fossorial Scincid Lizards?

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Biological Journal of the Linnean Society, 2020, XX, 1–10. With 2 figures.

Trade-offs between burrowing and biting force in fossorial scincid lizards?

MARGOT LE GUILLOUX1, AURÉLIEN MIRALLES2, JOHN MEASEY3, BIEKE VANHOOYDONCK4, JAMES C. O’REILLY5, AURÉLIEN LOWIE6, and

,

ANTHONY HERREL1,4,6,

*

1UMR 7179 C.N.R.S/M.N.H.N., Département Adaptations du Vivant, Bâtiment d’Anatomie Comparée, 55 rue Buffon, 75005, Paris, France 2Institut de Systématique, Evolution, Biodiversité, (UMR 7205 Muséum national d’Histoire naturelle, CNRS UPMC EPHE, Sorbonne Universités), CP30, 25 rue Cuvier 75005, Paris, France 3Centre for Invasion Biology, Department of Botany & Zoology, Stellenbosch University, Private Bag X1, 7602 Matieland, Stellenbosch, South Africa 4Deparment of Biology, University of Antwerp, Universiteitsplein 1, B2610 Antwerpen, Belgium 5Department of Biomedical Sciences, Ohio University, Cleveland Campus, SPS-334C, Cleveland, 45701 Ohio, USA 6Department of Biology, Evolutionary Morphology of Vertebrates, Ghent University, K.L. Ledeganckstraat 35, 9000 Ghent, Belgium

Received 27 December 2019; revised 17 February 2020; accepted for publication 19 February 2020

Trade-offs are thought to be important in constraining evolutionary divergence as they may limit phenotypic diversification. The cranial system plays a vital role in many functions including defensive, territorial, predatory and feeding behaviours in addition to housing the brain and sensory systems. Consequently, the morphology of the cranial system is affected by a combination of selective pressures that may induce functional trade-offs. Limbless, head-first burrowers are thought to be constrained in their cranial morphology as narrow heads may provide a functional advantage for burrowing. However, having a wide and large head is likely beneficial in terms of bite performance. We used 15 skink species to test for the existence of trade-offs between maximal push and bite forces, and explored the patterns of covariation between external head and body morphology and performance. Our data show that there is no evidence of a trade-off between bite and burrowing in terms of maximal force. Species that generate high push forces also generate high bite forces. Our data also show that overall head size covaries with both performance traits. However, future studies exploring trade-offs between force and speed or the energetic cost of burrowing may reveal other trade-offs.

ADDITIONAL KEYWORDS: covariation – cranial system – divergence – head-first burrowers – morphology – skink.

anatomical structures (Arnold,1992;Van Damme et al., 2002, 2003). Previous studies have further suggested that trade-offs may in some cases limit phenotypic variation by constraining evolutionary divergence

(Vanhooydonck et al., 2001; Levinton & Allen, 2005; Konuma & Chiba, 2007; Herrel et al., 2009). The

cranial system plays a vital role in many activities including defensive, territorial, predatory and feeding behaviours in addition to housing and protecting the brain and major sensory organs (Andrews et al., 1987;

Cooper & Vitt, 1993; Herrel et al., 2007; Kohlsdorf

INTRODUCTION

The phenotype of an organism reflects the selective pressures exerted by activities that are essential to its survival and its reproduction (Arnold, 1993). Sometimes, however, the functional demands exerted by different performance traits may result in tradeoffs. Indeed, functional trade-offs arise when different functions exert conflicting pressures on the same

*Corresponding author. E-mail: anthony.[email protected] © 2020 The Linnean Society of London, Biological Journal of the Linnean Society, 2020, XX, 1–10

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Burrowing and Biting in Scincid Lizards

et al., 2008; Dumont et al., 2016). Consequently, the

morphology of the cranial system is affected by these combined selective pressures which may potentially induce functional trade-offs. burrowing force. Consequently, trade-offs between these two performance traits, if present, are not mediated by direct functional conflict for the optimization of single functional trait. Rather, burrowers can be expected to have narrow heads for efficient soil penetration which may come at a cost of bite force.
The hypothetical trade-off between biting and borrowing performance is particularly interesting in limbless burrowing animals. These organisms are obligate head-first burrowers and the evolution of their morphology is thought to be constrained. Indeed, because the energy required for burrowing increases exponentially with body and head diameter (Navas et al., 2004), having a thin body and a narrow head may provide a functional advantage for burrowing.Yet, this is likely detrimental in terms of bite performance

(Verwaijen et al., 2002; Navas et al., 2004; Herrel & O’Reilly, 2006; Vanhooydonck et al., 2011; Baeckens

et al., 2017; Hohl et al., 2017). Maximum bite force has

been suggested to limit the type and size of food an

animal can eat (Herrel et al., 1999, 2001, 2008; Aguirre et al., 2003; McBrayer & Corbin, 2007; Edwards et al.,

2013). Consequently, a cranial form optimized for soil penetration may compromise the types of food an animal can eat and vice versa (Andrews et al., 1987;

Barros et al., 2011; Baeckens et al., 2017).

Here, we test for the presence of a trade-off between maximal bite force and the maximal push force in a range of burrowing and leaf-litter dwelling skinks. Scincid lizards represent an ideal study system as this family includes a variety of ecologies and morphologies with quadrupedal surface-dwelling species, epigeal serpentiform species with partially reduced front- and/ or hindlimbs, burrowing completely limbless species, and a plethora of intermediate forms (Pianka & Vitt, 2003;

Miralles et al., 2015; Wagner et al., 2018; Bergmann &

Morinaga, 2019). At least 15 independent evolutions of a serpentiformbodyformhavetakenplacewithinthegroup (Benesh & Withers, 2002; Miralles et al., 2012) allowing for a robust framework to test for associations between life-style, performance and morphology. Consequently, we also explore the patterns of covariation between head and body morphology and the two performance traits studied here (bite force and push force).
Burrowing is a complex behaviour that remains rather poorly understood in limbless head-first burrowing vertebrates (but see for example Gaymer

(1971); Gans (1973); O’Reilly et al. (1997); Teodecki et al. (1998); Navas et al. (2004); De Schepper et al. (2005)).

The maximal push force that an animal can generate is likely important as it may allow an animal to penetrate a greater variety of soil types, and thus expand its resource base in terms of potential habitat and food resources. As limbless species burrow by recruiting muscles along the long axis of the body (Rieppel, 1981;

Navas et al., 2004; Vanhooydonck et al., 2011; Hohl

et al., 2017), the diameter and size of the body should be related to the maximal push force it can generate. However, to facilitate soil penetration the width of the head should rather be narrow as this optimizes the pressure for a given force (e.g. Measey & Herrel, 2006;

Herrel & Measey, 2010; Barros et al., 2011). Moreover,

the speed by which it can penetrate the soil (and not only the force generated) is also likely a factor significantly contributing to the burrowing performance (Ducey

et al., 1993; Teodecki et al., 1998; Vanhooydonck et al.,

2011). Although few quantitative data exist, a previous study suggested the presence of a trade-off between bite force and burrowing speed in a limbless skink, Acontias percevali, mediated by the conflicting demands on head dimensions (Vanhooydonck et al., 2011). However, whether this is more generally the case and whether trade-offs also exist between bite force and push force remains unknown. As burrowing force is dependent on the axial musculature, different anatomical traits are responsible for the generation of bite force vs.

MATERIALS AND METHODS

AnimAls

Morphological measurements were performed on 180 individuals and performance measurements were obtained for 171 individuals for bite forces and 159 for push forces across 14 different species (Table 1).Animals were sampled between 2000 and 2017. The specimens were adults of often unknown sex. Data were collected in situ in the field or in the lab for species that were obtained through the pet trade. An additional five individuals of the species Pygomeles braconnieri from the collections of the National Museum of Natural History in Paris were used for morphological measurements.

morphometrics

Each individual was weighed using an electronic balance (Ohaus, 0.1 g). Head length, head width, head depth and lower jaw length were measured using a digital caliper (Mitutoyo, 0.01 mm) as described previously (Herrel and Holanova, 2008). The snoutvent length was measured by stretching the animals along a ruler ( 1 mm).

mAximAl push force

Maximalpushforcesweremeasuredinthefieldorinthe lab following the protocol described in Vanhooydonck

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M. LE GUILLOUX ET AL.

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© 2020 The Linnean Society of London, Biological Journal of the Linnean Society, 2020, XX, 1–10

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Burrowing and Biting in Scincid Lizards

Figure 1. Phylogeny used in the analyses modified from Pyron et al. (2013).

et al. (2011). Measurements of peak push forces during

mAximAl bite force

burrowing were made using a custom piezoelectric
Maximal bite forces were measured in the field or

the lab following Herrel et al. (1999). In brief, in vivo bite forces were measured using an isometric Kistler force transducer (type 9203, Kistler Inc.) mounted on a purpose-built holder and connected to a Kistler charge amplifier (type 5058 A, Kistler Inc.). Biting causes the upper plate to pivot around the fulcrum, and thus pull is exerted on the transducer. Capture of the animals typically resulted in a characteristic threat response where the jaws are opened maximally. The free end of the holder was then placed between the jaws of the animal, which immediately resulted in fierce and prolonged biting. The gape angle ( 30 °) and the place of application of the bite force was standardized with animals always biting at the tips of the jaws. Measurements were repeated five times for each animal and the maximum value recorded was considered to be the maximal bite force for that animal. force platform (Kistler Squirrel force plate, 0.1 N, Kistler Inc., Switzerland). The force platform was positioned on a custom-designed metal base and connected to a charge amplifier (Kistler Charge Amplifier type 9865, Kistler Inc.).A Perspex block with 1 cm-deep holes of different diameters was mounted on the force plate, level with the front edge. One of the holes was loosely filled with soil from the container of the animal that was tested. A Perspex tunnel with a diameter similar to the maximal body diameter of the test animal was mounted on the metal base in front of (but not touching) the force plate, and aligned with the soil-filled hole in the Perspex block. First, a skink was introduced into the tunnel and allowed to move through it until reaching the soil-filled chamber. Next, the animal was stimulated to burrow into the soil by tapping the end of the tail sticking out of the tunnel, or by prodding the animal inside the tunnel with the blunt end of a thin wooden stick. Forces were recorded during a 60 s recording session at 500 Hz, and three trials were performed for each individual, with at least 1 h between trials. Forces were recorded in three dimensions using the Bioware software (Kistler Inc.). For each individual we then extracted the highest peak resultant force across all trials as an indicator of that animal’s maximal push force.

AnAlyses

  • Morphometric and force data were Log10
  • -

transformed before analysis to ensure normality and homoscedasticity. All analyses were performed in R (v.3.4.0) while taking into account the phylogenetic relationships among species. The phylogenetic framework

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M. LE GUILLOUX ET AL.

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Figure 2. Phylogenetic regressions of species averages of maximal bite force against maximal push force. (A) Absolute push force is significantly correlated with absolute bite force. (B) Residual bite force is significantly and positively correlated with residual push force when correcting for overall snout-vent length. (C) Residual bite force is no longer related to variation in residual push force when corrected for variation in body mass.

© 2020 The Linnean Society of London, Biological Journal of the Linnean Society, 2020, XX, 1–10

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Burrowing and Biting in Scincid Lizards

Figure 3. Covariance between morphological and performance data. (A) Phylogenetic two-block partial least square analysis between morphology and performance illustrating that more robust species (i.e. with greater body mass and bigger heads) produce greater push and bite forces. (B) Two-block partial least square analysis on the force and morphological data corrected for variation in snout-vent length. Species with relatively higher mass, heads size and body diameter produce greater bite and push forces. (C) Two-block partial least square analysis on the force and morphological data corrected for variation in body mass showing that species that are relatively more elongate (higher snout-vent length for their mass) show relatively higher push forces. In contrast, species that are stockier, less elongate and with bigger heads produce relatively higher bite forces.

used was obtained from the molecular data set of Pyron et al. (2013); Fig. 1. This required the reconstruction of a phylogeny by pruning the tree to include only species for which we had performance and morphological data. First, a phylogenetic generalized least squares (PGLS) regression was

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M. LE GUILLOUX ET AL.

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run in R (function gls with corBrownian in Phytools (Revell, 2012)) with the mean maximal push force per species against mean maximal bite forces to test for the presence of a trade-off between bite and push force. To test for co-variation between morphology and performance we ran a phylogenetic two-block partial least squares analysis (φ2bPLS) using the function ‘phylo.integration’ implemented in ‘geomorph’ in R (Adams et al., 2013). This method quantifies the degree of association between two data matrices. It is a descriptive multivariate analysis robust to multicollinearity between variables and therefore suitable for the use of morphometric and performance variables. The analysis generates axes that explain the covariance between the two data matrices. As body size simultaneously impacts head and body dimensions and forces, we ran PGLS analyses with snout-vent length (SVL) or body mass as our predictor and morphometric and performance traits as our independent variables and extracted the unstandardized residuals. Next, we used a Pearson correlation test to explore the existence of a tradeoff between our two residual performance traits independent of variation in overall body size. Finally, we ran a two-block partial least squares analyses (2BPLS) on the residual data to explore patterns of covariation between morphology and performance independent of variation in overall size using‘two.b.pls’ script of the ‘geomorph’ package (Adams et al., 2013), and ‘pls2B’ script of the ‘Morpho’ package (Schlager, 2013) in R. appear to have relatively large bite forces but low push forces.

covAriAtion between morphology And performAnce

The phylogenetic 2BPLS analysis was significant (rPLS = 0.983, P = 0.001). Heavier, larger and wider animals with longer, taller and wider heads produce larger forces (Fig. 3A). The 2BPLS analysis run on the size-corrected data was also significant (rPLS = 0.969, P < 0.001; Fig. 3B).This analysis indicates that the maximal push force and the maximal bite force co-vary, both principally with overall head and body robustness, with animals that are more elongate producing relatively lower push and bite forces. An analysis on the size corrected data using body mass was also significant (rPLS = 0.73, P = 0.012), yet showed a different pattern. This analysis shows that more elongate animals produce relatively higher push forces whereas the more robust limbed species with wide heads and bodies produce low push but high bite forces for their body mass (Fig. 3C).

DISCUSSION

Our results, based on a broad range of burrowing and leaf-litter dwelling skinks, show that there is no direct trade-off between bite force and burrowing force in this group. Species that produce strong push forces are also those which produce strong bite forces in both absolute and relative terms. These results suggest that whereas bite force may trade-off with burrowing

speed (Teodecki et al., 1998; Vanhooydonck et al., 2011)

this may not be the case for push force. Indeed, the same traits that favour increased bite force (large, robust heads and wide bodies) also appear to favour high push forces, at least in absolute terms. This makes intuitive sense as the muscles used to generate both bite and push force are positioned to the lateral side of the body in scincid lizards (Huyghe et al., 2009; Vanhooydonck et al., 2011). For example, the external adductor muscle is one of the largest contributors to overall bite force generation and lies external to the mandible (Vanhooydonck et al., 2011). Similarly, the iliocostalis and longissimus muscles that generate the push forces measured are also positioned laterally to the vertebral column. Consequently, wider heads and wider bodies should induce an increase in both absolute bite and push force. However, whereas these traits may indeed favour absolute force, the speed by which animals can penetrate the soil may be negatively impacted by the presence of wider heads

and bodies (Teodecki et al., 1998; Vanhooydonck et al.,

RESULTS

trAde-offs between bite force And push force

The linear regression taking into account phylogeny shows that maximum push force is positively correlated with maximal bite force (PGLS: r = 1.36, P < 0.001), suggesting that species that produce strong push forces are also those who produce strong bite forces (Fig. 2A). Analyses performed on the snoutvent length corrected data show a similar result (r = 0.89, P < 0.001) with animals that bite harder for a given size also being better pushers (Fig. 2B). However, when correcting force measurements for body mass, residual bite force was no longer correlated with residual push force (r = -0.63, P = 0.053; Fig. 2C). An inspection of the plot (Fig. 2C) suggests that more elongate and smaller species like Typhlosaurus

vermis and T. l omiae as well as Acontias litoralis

and A. kgalagadi tend to have relatively higher push forces whereas the stockier, larger species like

Scincus scincus, A. meleagris, and Mochlus sundevalli

© 2020 The Linnean Society of London, Biological Journal of the Linnean Society, 2020, XX, 1–10

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Burrowing and Biting in Scincid Lizards
2011). Moreover, the energetic cost of burrowing may also be negatively impacted by these traits (Navas et al., 2004). However, whereas the limbed species are the best biters in absolute terms and when corrected for body mass the more elongate specialized burrowers

(e.g. Acontias, Scelotes, Typhlosaurus) actually produce

greater push forces. Indeed, the specialized head-first

limbless burrowers like Acontias or Typhlosaurus

have higher push forces for their mass despite being more elongate and less robust. Moreover, individuals of these species were usually captured at deeper soil depths and in less sandy soils (A. Herrel & J. Measey, pers. obs.) suggesting that they are overall better at burrowing. Thus, rather than absolute push force relative push force may be the principal driver of burrowing specialization. However, given the paucity of quantitative data on soil hardness and burrow depth in fossorial animals a quantitative analysis of these patterns is not possible. Additionally, it would be of interest to gather similar data on other groups of fossorial skinks from different radiations (Australia, Madagascar, the Philippines, see Wagner et al. (2018);

Morinaga & Bergmann, 2020) as the data in the

present study are strongly biased towards African skinks, especially the Acontinae.

Vanhooydonck et al., 2011; Hohl et al., 2017) as also

suggested by our data. Indeed, more elongate species of the genera Typhlosaurus as well as the more elongate

Acontias species (A. litoralis, A. kgalagadi) showed

high push forces yet low bite forces (Fig. 3C). It would consequently be of interest to perform finer scale analyses of head shape or skull shape using geometric morphometric approaches. This may also allow teasing apart of differences between the back and the front of the skull likely impacted by constraints on biting vs.

burrowing (Cornette et al., 2015).

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  • Comparative Study of the Osteology and Locomotion of Some Reptilian Species

    Comparative Study of the Osteology and Locomotion of Some Reptilian Species

    International Journal Of Biology and Biological Sciences Vol. 2(3), pp. 040-058, March 2013 Available online at http://academeresearchjournals.org/journal/ijbbs ISSN 2327-3062 ©2013 Academe Research Journals Full Length Research Paper Comparative study of the osteology and locomotion of some reptilian species Ahlam M. El-Bakry2*, Ahmed M. Abdeen1 and Rasha E. Abo-Eleneen2 1Department of Zoology, Faculty of Science, Mansoura University, Egypt. 2 Department of Zoology, Faculty of Science, Beni-Suef University, Beni-Suef, Egypt. Accepted 31 January, 2013 The aim of this study is to show the osteological characters of the fore- and hind-limbs and the locomotion features in some reptilian species: Laudakia stellio, Hemidactylus turcicus, Acanthodatylus scutellatus, Chalcides ocellatus, Chamaeleo chamaeleon, collected from different localities from Egypt desert and Varanus griseus from lake Nassir in Egypt. In the studied species, the fore- and hind-feet show wide range of variations and modifications as they play very important roles in the process of jumping, climbing and digging which suit their habitats and their mode of life. The skeletal elements of the hand and foot exhibit several features reflecting the specialized methods of locomotion, and are related to the remarkable adaptations. Locomotion is a fundamental skill for animals. The animals of the present studies can take various forms including swimming, walking as well as some more idiosyncratic gaits such as hopping and burrowing. Key words: Lizards, osteology, limbs, locomotion. INTRODUCTION In vertebrates, the appendicular skeleton provides (Robinson, 1975). In the markedly asymmetrical foot of leverage for locomotion and support on land (Alexander, most lizards, digits one to four of the pes are essentially 1994).
  • An Etymological Review of the Lizards of Iran: Families Lacertidae, Scincidae, Uromastycidae, Varanidae

    An Etymological Review of the Lizards of Iran: Families Lacertidae, Scincidae, Uromastycidae, Varanidae

    International Journal of Animal and Veterinary Advances 3(5): 322-329, 2011 ISSN: 2041-2908 © Maxwell Scientific Organization, 2011 Submitted: July 28, 2011 Accepted: September 25, 2011 Published: October 15, 2011 An Etymological Review of the Lizards of Iran: Families Lacertidae, Scincidae, Uromastycidae, Varanidae 1Peyman Mikaili and 2Jalal Shayegh 1Department of Pharmacology, School of Medicine, Urmia University of Medical Sciences, Urmia, Iran 2Department of Veterinary Medicine, Faculty of Agriculture and Veterinary, Shabestar Branch, Islamic Azad University, Shabestar, Iran Abstract: The etymology of the reptiles, especially the lizards of Iran has not been completely presented in other published works. Iran is a very active geographic area for any animals, and more especially for lizards, due to its wide range deserts and ecology. We have attempted to ascertain, as much as possible, the construction of the Latin binomials of all Iranian lizard species. We believe that a review of these names is instructive, not only in codifying many aspects of the biology of the lizards, but in presenting a historical overview of collectors and taxonomic work in Iran and Middle East region. We have listed all recorded lizards of Iran according to the order of the scientific names in the book of Anderson, The Lizards of Iran. All lizard species and types have been grouped under their proper Families, and then they have been alphabetically ordered based on their scientific binominal nomenclature. We also examined numerous published works in addition to those included in the original papers presenting each binomial. Key words: Etymology, genera, iran, lizards, Middle East, species, taxonomy. INTRODUCTION comprising the fauna of Iran, including Field guide to the reptiles of Iran, (Vol.
  • The Terrestrial Mammals, Reptiles and Amphibians of the Uae – Species List and Status Report

    The Terrestrial Mammals, Reptiles and Amphibians of the Uae – Species List and Status Report

    THE TERRESTRIAL MAMMALS, REPTILES AND AMPHIBIANS OF THE UAE – SPECIES LIST AND STATUS REPORT January 2005 TERRESTRIAL ENVIRONMENT RESEARCH CENTRE ENVIRONMENTAL RESEARCH & WILDLIFE DEVELOPMENT AGENCY P.O. Box 45553 Abu Dhabi DOCUMENT ISSUE SHEET Project Number: 03-31-0001 Project Title: Abu Dhabi Baseline Survey Name Signature Date Drew, C.R. Al Dhaheri, S.S. Prepared by: Barcelo, I. Tourenq, C. Submitted by: Drew, C.R. Approved by: Newby, J. Authorized for Issue by: Issue Status: Final Recommended Circulation: Internal and external File Reference Number: 03-31-0001/WSM/TP007 Drew, C.R.// Al Dhaheri, S.S.// Barcelo, I.// Tourenq, C.//Al Team Members Hemeri, A.A. DOCUMENT REVISION SHEET Revision No. Date Affected Date of By pages Change V2.1 30/11/03 All 29/11/03 CRD020 V2.2 18/9/04 6 18/9/04 CRD020 V2.3 24/10/04 4 & 5 24/10/04 CRD020 V2.4 24/11/04 4, 7, 14 27/11/04 CRD020 V2.5 08/01/05 1,4,11,15,16 08/01/05 CJT207 Table of Contents Table of Contents ________________________________________________________________________________ 3 Part 1 The Mammals of The UAE____________________________________________________________________ 4 1. Carnivores (Order Carnivora) ______________________________________________________________ 5 a. Cats (Family Felidae)___________________________________________________________________ 5 b. Dogs (Family Canidae) __________________________________________________________________ 5 c. Hyaenas (Family Hyaenidae) _____________________________________________________________ 5 d. Weasels (Family Mustelidae) _____________________________________________________________
  • Felis Margarita, Sand Cat

    Felis Margarita, Sand Cat

    The IUCN Red List of Threatened Species™ ISSN 2307-8235 (online) IUCN 2008: T8541A50651884 Felis margarita, Sand Cat Assessment by: Sliwa, A., Ghadirian, T., Appel, A., Banfield, L., Sher Shah, M. & Wacher, T. View on www.iucnredlist.org Citation: Sliwa, A., Ghadirian, T., Appel, A., Banfield, L., Sher Shah, M. & Wacher, T. 2016. Felis margarita. The IUCN Red List of Threatened Species 2016: e.T8541A50651884. http://dx.doi.org/10.2305/IUCN.UK.2016-2.RLTS.T8541A50651884.en Copyright: © 2016 International Union for Conservation of Nature and Natural Resources Reproduction of this publication for educational or other non-commercial purposes is authorized without prior written permission from the copyright holder provided the source is fully acknowledged. Reproduction of this publication for resale, reposting or other commercial purposes is prohibited without prior written permission from the copyright holder. For further details see Terms of Use. The IUCN Red List of Threatened Species™ is produced and managed by the IUCN Global Species Programme, the IUCN Species Survival Commission (SSC) and The IUCN Red List Partnership. The IUCN Red List Partners are: BirdLife International; Botanic Gardens Conservation International; Conservation International; Microsoft; NatureServe; Royal Botanic Gardens, Kew; Sapienza University of Rome; Texas A&M University; Wildscreen; and Zoological Society of London. If you see any errors or have any questions or suggestions on what is shown in this document, please provide us with feedback so that we can correct or extend the information provided. THE IUCN RED LIST OF THREATENED SPECIES™ Taxonomy Kingdom Phylum Class Order Family Animalia Chordata Mammalia Carnivora Felidae Taxon Name: Felis margarita Loche, 1858 Regional Assessments: • Mediterranean Common Name(s): • English: Sand Cat, Sand Dune Cat • French: Chat des sables • Spanish: Gato de las Arenas, Gato del Sahara Taxonomic Notes: Taxonomy is currently under review by the IUCN SSC Cat Specialist Group (2014).
  • "Mimicking the Abrasion Resistant Sandfish Epidermis"

    "Mimicking the Abrasion Resistant Sandfish Epidermis"

    "Mimicking the abrasion resistant sandfish epidermis" Von der Fakultät für Mathematik, Informatik und Naturwissenschaften der RWTH Aachen University zur Erlangung des akademischen Grades eines Doktors der Naturwissenschaften genehmigte Dissertation. vorgelegt von Boštjan Vihar, M.Sc. Aus Maribor, Slowenien Berichter: Universitätsprofessor Prof. Dr. Peter Bräunig Universitätsprofessor Prof. Dr. Werner Baumgartner Tag der mündlichen Prüfung: 7. September 2015 Diese Dissertation ist auf den Internetseiten der Universitätsbibliothek online verfügbar. "L'essentiel est invisible pour les yeux." Antoine de Saint-Exup´ery Abstract The sand inhabiting skink Scincus scincus has due to its sand swimming behaviour gained some attention by the research community in recent years. The sand swimming creates ample stress on the outer layers of its skin, generating the prospect of wear. The skin itself is however very resilient towards abrasion and even facilitates smoother movement by reducing friction against the sharp sand particles. Past research has concentrated much on the characterization of structure and surface properties of the skin, however, a complete understanding of its properties is lacking up to date. In this work, a review of known data is done in addition to new measurements concerning surface properties, composition, ultrastructure and mechanics, but also other feats such as optical properties. Replication of "anti-adhesive" properties on existing technical materials was tested along with the development of new techniques for such endeavour. Chemical and structural anal- ysis shows the epidermis of the sandfish is very homogeneous and is mainly composed of compact bundles of β keratins, which are possibly interlinked by a matrix composed of α keratins. Strong glycosylation of the keratins was shown, confirming previous studies, and five glycan structures were observed in the analysis.
  • Patterns of Species Richness, Endemism and Environmental Gradients of African Reptiles

    Patterns of Species Richness, Endemism and Environmental Gradients of African Reptiles

    Journal of Biogeography (J. Biogeogr.) (2016) ORIGINAL Patterns of species richness, endemism ARTICLE and environmental gradients of African reptiles Amir Lewin1*, Anat Feldman1, Aaron M. Bauer2, Jonathan Belmaker1, Donald G. Broadley3†, Laurent Chirio4, Yuval Itescu1, Matthew LeBreton5, Erez Maza1, Danny Meirte6, Zoltan T. Nagy7, Maria Novosolov1, Uri Roll8, 1 9 1 1 Oliver Tallowin , Jean-Francßois Trape , Enav Vidan and Shai Meiri 1Department of Zoology, Tel Aviv University, ABSTRACT 6997801 Tel Aviv, Israel, 2Department of Aim To map and assess the richness patterns of reptiles (and included groups: Biology, Villanova University, Villanova PA 3 amphisbaenians, crocodiles, lizards, snakes and turtles) in Africa, quantify the 19085, USA, Natural History Museum of Zimbabwe, PO Box 240, Bulawayo, overlap in species richness of reptiles (and included groups) with the other ter- Zimbabwe, 4Museum National d’Histoire restrial vertebrate classes, investigate the environmental correlates underlying Naturelle, Department Systematique et these patterns, and evaluate the role of range size on richness patterns. Evolution (Reptiles), ISYEB (Institut Location Africa. Systematique, Evolution, Biodiversite, UMR 7205 CNRS/EPHE/MNHN), Paris, France, Methods We assembled a data set of distributions of all African reptile spe- 5Mosaic, (Environment, Health, Data, cies. We tested the spatial congruence of reptile richness with that of amphib- Technology), BP 35322 Yaounde, Cameroon, ians, birds and mammals. We further tested the relative importance of 6Department of African Biology, Royal temperature, precipitation, elevation range and net primary productivity for Museum for Central Africa, 3080 Tervuren, species richness over two spatial scales (ecoregions and 1° grids). We arranged Belgium, 7Royal Belgian Institute of Natural reptile and vertebrate groups into range-size quartiles in order to evaluate the Sciences, OD Taxonomy and Phylogeny, role of range size in producing richness patterns.